q
I
•
and each infected egg has therefore to be carefully eliminated.
Even by observing these rules of caution, more strings of eggs
died and had to be thrown into the bucket than Monsieur
Boulenger presumably used in his experiments, until in the end
I managed to succeed with a very few eggs from a very few
strings.' 18
Boulenger quotes this reply only to accuse Kammerer that he
invented this excuse when cornered: 19
Why, it may be asked, was all this not mentioned at first,
instead of letting the reader believe that the embryos unqer-
went the whole of their development without any interven-
tion on the part of the experimentator.
From this sample of the levity with which Kammerer
relates his experiments, is it surprising if some of his state-
ments should be challenged by those who, like myself, do
not place implicit confidence in them?
But the levity-if that is the right word-is on Boulenger's
side. It is simply untrue to say that Kammerer did not, from the
very beginning of his experiments, explicitly point out the pre-
cariousness of Alytes eggs submerged in water. He did so in his
first paper on the subject, published in 190620 (Kammerer's
italics) :
Before describing the next experiment, I must give some
technical indications for the correct preservation of the eggs
destined to develop without the paternal care of the male.
I deposit the balls of egg-strings in glass cups on fine river
sand, which has to be sterilised before use to eliminate
fungus germs. According to the nature of the experiment,
the sand is kept moist or wet in varying degrees. In a
humid environment, the cup is closed with a well-fitting
lid. If the eggs are made to develop in darkness, they are
covered with sterilised blotting paper which is moistened,
instead of the sand. Earth and moss, though natural media,
are to be avoided, because otherwise most of the eggs are
attacked by mould. Those string-balls which are to have
daily baths are extracted-always at the same time of day-
by means of a horn spoon from the sand bed and placed for
five minutes into a cup of water. In the case of eggs reared
in darkness, this manipulation is carried out in a darkroom.
In spite of all these precautions, one is liable to lose a fairly
155
high percentage of the eggs through withering or the pro-
liferation of mould fungus so that, to be able to complete
an experiment, one has to start with substantial material.
This is particularly the case in Experiment NO.4 to be
presently described: The Maturation of Alytes Eggs
Deprived of Paternal Care, in Water . ...
And he did warn again, in his second paper on the subject,
published in 1909:21
It should be repeated once more that the Alytes eggs kept
in water are so vulnerable to begin with that on some
strings no eggs at all reach maturity, on other strings only
3 per cent to 5 per cent.
It is indeed astonishing that Boulenger has not only failed to
take these warnings into account, but denied that they had
ever been uttered. Even more astonishing: the first quote above
ends on the same page of the 1906 paper-page 69-to which
Boulenger referred as saying that all Alytes mated on the same
day. He thus could not have overlooked the technical instruc-
tions whose existence he denied. No wonder he failed to repeat
Kammerer's experiments; but this can hardly be construed as
an 'elaborate and destructive criticism'.
At the end of his article Boulenger repeated his criticism
about the position of the nuptial pad, quoting at length what
was said on the subject by Bateson-and yet once more Bate-
son's letter to Kammerer in 1910, and the latter's failure to
produce a specimen.
In spite of having 'often wished to visit the Vivarium',
Boulenger never did, and thus never had an opportunity to
acquaint himself, by first-hand experience, with Kammerer's
special techniques-or with the distance between Hiitteldorf
and Vienna. After the First World War he fades out of the
picture. Kammerer's 'reply to Boulenger' (which I shall quote
in a moment) was published in his long 1919 paper which
Boulenger had 'not put himself out to procure'; he never
answered it.
His son, E. G. Boulenger, did, however, visit the Vienna
Institute in 1922. Although he too had been sceptical about
Kammerer's claims, he seems to have changed his mind and
came back with the conviction that 'if we still disbelieve, we
must assume that Przibram is a dishonest person'. He attended
Kammerer's Cambridge lecture and demonstration, but did not
take part in the discussion, nor in the controversy in. Nature.""
The last document in the Boulenger controversy IS Kammer-
er's reply in his concluding 1919 paper on Alytes. It occupies
forty-five pages in the Archiv fur Entwicklungsmechanik, of
which eight are devoted to replies to his critics-Boulenger,
Bateson and Bauer. I shall quote the reply to Boulenger almost
in full, since it sums up several vital aspects of the controversy
(Kammerer's italics):
Reply to Boulenger
I have a strong aversion against polemics. That is the reason
why I have not answered be!ore vari?us .attacks o ~ the
reliability of my results and theIr theoretlcal mterpretatlOns.
I postponed my reply until new results, and not just t ~ e
desire to answer back, would justify it. As far as Alytes 1S
concerned, this is now the case.
Boulenger is one of the few who have attempted to check
my results by experiment before condemning them. He
took the freshly laid and fertilised eggs from two (two!)
males after observing their copulatory acts, and threw the
eggs into the water of a trough (,abreuvoir')t in which he
had seen Alytes tadpoles; he therefore thought the con-
ditions ideal for his experiment.
Boulenger was lucky for apparently his eggs developed
normally for as long as 5-6 days before they died. For, as I
have already explained in 1914 in a short paper in reply to
Boulenger, I would have expected that under these 'natural'
conditions the eggs would have been totally infested with
fungi. And already in 1906 I reported my own difficulties
in keeping the 'water' -eggs free from saprolegniacea [a
group of fungi], and recommended maintenance under as
sterile conditions as possible. Boulenger passes this passage
in silence ....
Boulenger, who by the nature of his work is understand-
"" Boulenger, jr. (who died in 1946), seems to have been a person of
great charm and humour. He travelled a lot, collecting specimens,
always by sea, and mostly on foreign boats. A friend of mine, who
knew him well, once asked him the reason for this preference, and got
the startling reply: 'They have none of this nonsense about women and
children first ... .'
t Kammerer does not realise that 'abreuvoir' may also mean a small
pond; but the point is irrelevant.
157
ably unacquainted with experimental methods, did not
realise what every physiologist would take for granted:
that even boiled and artificially ventilated water is not im-
mune against invasion by mould germs; each affected egg
must be carefully removed, cut away from the ball (of the
tangled strings) with a pair of fine scissors, and the sterilisa-
tion of the whole container must then be repeated. In spite
of these precautions I had to throwaway many more balls,
after the embryos they contained had died, than Boulenger
has ever handled. Later on, however, the results improve:
in successive generations the mortality of 'water' -eggs is
hardly greater than that of other ·anura who depose their
eggs normally in water. But the road that leads to that stage
is long. A few field observations and a basin fined with
water from a puddle provide no short-cut.
Breeding Alytes from 'water' -eggs is the precondition
for the appearance, in later generations, of nuptial pads
during the mating season. And since Boulenger failed in
this form of breeding, he doubts in the same breath the
existence of the pads. Yet he has an added argument for his
disbelief: that in 1909 I described and drew the pad only
on one (the innermost) finger. Boulenger however saw that
during amplexus [the mating embrace], in contradistinction
to frogs, not one but two fingers enter into contact with the
female's pubic region. This statement is derived from
observing a single (!) copulating Alytes pair, which
Boulenger picked up. He concludes: if pads did show at all
on my water-mating specimens, they would have had to
appear, logically on two fingers and not on one.
How can Boulenger state, with such assurance, that the
position of these very timorous animals was not displaced
when he took them in his hand-and without picking them
up, they cannot be inspected. How can he pretend to know
-assuming that his description is correct-that both
fingers were used with equal strength? In view of the un-
doubted variability of the mating postures of Alytes (cf.
the observations of Dahne), how can Boulenger generalise
from his one single observation, and moreover extend his
generalisation from land-mating to water-mating Alytes? I
have certainly watched more Alytes copulations-on land
and in water-than Boulenger's three cases; and I regard it
as important that I watched them at close quarters in the
158
terrarium-and not in the light of an electric torch in the
gutter of a village street or in the clefts of a stone in a ruin.
Yet I would not dare to make any generalised statement re-
garding the action of the male's fingers (which during
copulation are hidden under the female); nor as to the
number and position of the fingers which exert friction on
the female's skin. It is even possible that the variability and
extension of the nuptial pads, as apparent in the material
presented in this paper, corresponds to as many modifica-
tions in the positions of the embrace.
Boulenger can, in view of the evidence, hardly doubt any
longer the existence of the pads; in view of their variability
he will perhaps also realise that deductions from ethological
observations to morphological features require much
greater caution than he has shown.
Boulenger, as already said, did not reply.
As far as the literature indicates, Boulenger and Bateson were
the only zoologists who made the attempt to repeat Kammerer's
experiments with Alytes. Bateson did not publish his results.
The texts I have quoted explain why Boulenger failed; and
they indicate that if Kammerer's experiments were regarded as
unrepeatable and therefore suspect, the blame lies on those
who tried to repeat them with inadequate methods.
159
APPENDIX 4
The Location of the
Pads
It will be remembered that after the meeting at the Linnean
Society in 1923, Bateson published (Nature, June 2) his hither-
to most violent attack on Kammerer. Among the various points
which Bateson raised, the principal was that the dark mark on
the skin of the critical specimen of Alytes was 'in the wrong
place', namely, across the palm (Bateson's italics):
I direct attention first to the fact that the structure shown
did not look like a real Brunftschwiele. Next, I lay stress
on its extraordinary position. It was in the wrong place.
Commenting on the evidence, I pointed this out. In the
embrace of Batrachians the palms of the hand of the male
are not in contact with the female [i.e. the male turns the
backs of its hands to a certain extent inwards]. To show
how the hands are placed, I send a photograph (Figure 1) of
a pair of Rana agilis, killed and preserved while coupled.
Clearly the rugosities, to be effective, must be on the backs
and radial sides of the digits, round the base of the thumb,
as in our common frog, on the inner sides of the forearms,
or in certain other positions, but not on the palms of the
hands. There are, of course, minor variations, in correspon-
dence with which the positions of the rugosities differ. But
on the palm of Alytes they would be as unexpected as a
growth of hair on the palm of a man.
To this MacBride replied on June 23. Regarding the texture
of the pad, he pointed out that Bateson had omitted any refer-
160
ence to the microscope sections which Kammerer had shown;
and that the pad, in a Rana, too, 'looks like a simple patch of
pigment, and passing my finger over it, I could not detect the
capillae by feeling'. As to the position of the pad, MacBride
quoted Boulenger's description of the different positions of the
pads in different species-positions including regions which
never get into contact with the female.
Bateson replied very briefly on June 30, avoiding all issues
except for pointing out that when Boulenger spoke of the 'inner'
side of fingers showing the pad, he meant the radial, not the
palmar, side.
MacBride replied on July 21 that this was quite true, 'but the
callosity on the radial edge of the finger involves the palmar
surface also, as Dr. Bateson may convince himself by inspecting
Boulenger's figures, and as, indeed, is demonstrated to every
student when he is shown the nuptial callosity of the male Rana'.
Bateson did not reply.
On August 18 Kammerer, back from the U.S.A., replied
himself to Bateson's attack. On the crucial question he wrote:
It is incorrect to say that the black colour is restricted to
the palmar aspect. (Why should Mr. Bateson assert this
when he had not seen the dorsal aspect?) Actually the pads
extend to the dorsal aspect and are therefore not 'in the
wrong place'.
It is incorrect to say that the pad presents only 'a dark
uniform surface but no capillary or thorny structures'. I
send herewith an enlarged photograph in which 'rugosities'
can be seen on the edge of the pad with the naked eye.
In the same issue of Nature (August 18), Michael Perkins
also replied to Bateson. I shall quote him at some length be-
cause his letter went into more detail than any of the other
correspondents', and should really have settled the issue:
Dr. Bateson points to two details which make 'the appear-
ance quite unlike that of any natural Brunftschwielen':
first, that in Alytes there is a 'dark uniform surface ...
without the dotting or stippling so obvious in true Brunft-
schwielen'; secondly, that their position does not corres-
pond to that of the nuptial pads in Rana agilis.
Lataste's excellent drawings (Ann. Sci. Nat. (6), tom. 3,
pl. II, 1876) show that a uniform darkness of the outer
r6r
layer of the pad is a characteristic feature of the Disco-
glossidae (to which Alytes belongs) and distinguishes them
from other Anura. The fully developed pads of Bufo vul-
garis are also uniformly black, and I have recently found
that when such full hypertrophy of the outer epithelium is
inhibited, as occasionally happens from obscure causes, it
may be induced by making the male maintain a sexual em-
brace for a week or two ....
The pad of the Alytes 'water-breed' also resembles that
of the Discoglossid Bombinator in having a complete layer
of black pigment in the cutis vera which would further con-
tribute to the uniform dark appearance which Alytes so well
and characteristically shows ....
The epidermal spines are very obvious in the intact
specimen, as I have repeatedly seen both with lens and
binocular microscope, and as many others have witnessed
in my presence. Of course, they are practically impossible
to photograph on account of the glistening of a wet speci-
men, but a photograph at least makes clear what areas of
skin are affected. These include nearly the whole of the
palm, the radial surface of the inner metacarpal and part of
the first phalangeal joint of the thumb, and more or less of
the ventral and radial surfaces of the forearm, passing over
the dorso-radial margin of the inner carpal tubercle. The
Discoglossidae are remarkable for the very various positions
in which the histological features of Brunftschwielen may
manifest themselves, on the chin, belly, thighs, toes of the
feet even; in other words, they are not necessarily depen-
dent on contact with the female for their development. Dr.
H. Gadow has shown me his sketch of the nuptial pad in
Alytes cisternasii, Bosca., where it is d e v ~ p e d on the tip of
the thumb, extending on the palmar surface. Even in the
common toad I have frequently observed the nuptial
rugosity extending on to the palmar surface of the inner
carpal tubercle.
Questionable as it is to draw conclusions on anatomical
points by analogy from other animals, it is even more un-
safe to do so as regards their habits and postures; Alytes
does not belong even to the same suborder as Rana agilis.
Bateson replied to Kammerer and Perkins on September IS.
He completely ignored the factual contents of Perkins' letter,
162
I
j
f-
l.
I
which he dismissed in two sentences: 'Mr. Perkins states that
"the epidermal spines are very obvious in the intact specimen".
He is the only independent witness of those whose opinions
have reached me, who claims to have seen anything so definite.'
In his reply to Kammerer, Bateson simply repeated that the pad
was 'in the wrong place on the palm'; and that it 'did not look
like a nuptial pad .... What there may have been on the back of
the hand I do not know.' This was the letter which ended with
the offer of £25 for sending the Alytes back to London.
Przibram refused the offer in a letter to Bateson which I have
quoted on page 86. I have also quoted Bateson's reply to
Przibram: 'I would gladly now double my offer, etc'. But
Bateson's reply to Przibram, published in Nature on December
22, also contained a new, and quite unfounded, accusation
in the last but one sentence of the quotation which follows
(Bateson's italics):
In my last letter I explained how I missed making a proper
examination [of the specimen] here. Reports had varied,
and I drew the inference that the nature of the black marks
must be mainly a question of interpretation. Not until I
saw the toad at the Linnean meeting, with the unexpected
and misplaced development on the palm of the hand, did I
discover that there was anything so positive to examine. As
I thought over the incident it struck me as extraordinary
that this, the real peculiarity of the specimen-which, in-
deed, it was set up to display-had never been mentioned
by Dr. Kammerer. He left England immediately after the
meeting.
The phrase 'had never been mentioned by Dr. Kammerer'
seemed to imply that Kammerer was so embarrassed by the
shameful position of the pad on the palm that both in the past
and during his lecture he had passed it over in silence. But
Bateson's accusation happens to be demonstrably untrue. In
the text of Kammerer's lecture, which he gave both in Cam-
bridge and at the Linnean, published in Nature on May 12, we
read (my italics):
Of the many changes which gradually appear in this water
breed during the various stages of development-egg, larva,
and the metamorphosed animal, young and old-I will des-
cribe only one, the above-mentioned nuptial pad of the
16
3
male. At first it is confined to the innermost fingers, but in
subsequent breeding seasons it extends to the other fingers,
to the balls of the thumb, even to the underside of the
lower arm. Mter spreading, it exhibits an unexpected vari-
ability, both in the same individual and between one indi-
vidual and another. The variability in the same individual
is shown by the characters altering from year to year and in
the absence of symmetry between the right hand and the
left. In one specimen the dark pad extended to all the other
fingers and almost over the whole of the left hand.
But that is not all. By courtesy of Professor W. H. Thorpe
and the Cambridge Natural History Society, I have been able
to obtain the original typescript of Kammerer's lecture. There,
the last sentence just quoted reads as follows (my italics): 'I
have here a specimen in which the dark pad has extended, etc.'
(Obviously the Editor of Nature made one of the routine
corrections in the printed version of an oral delivery.)
Bateson's repeatedly expressed surprise at the 'unexpected
a ~ d misplaced development on the palm of the hand' is equally
dIfficult to understand. A few months earlier, the younger
Boulenger had, as we remember, visited the Vienna Institute
and then reported, in a letter to Bateson, that he had seen the
famous specimen of Alytes and that 'nearly the whole hand is
coloured black'. It is hard to believe that Bateson should have
forgotten this. Then why pretend to be so astonished by the
sight of the specimen-and why refuse to examine it, as others
did?
Bateson's letter was to have a distressing influence on
Przibram, but only four years later. Its immediate effect must have
been to incense even that saintly man by the humiliating offer
of first £25, then £50 for sending the specimen back to London,
so Przibram probably did not pay much attention to the insinua-
tions it contained. But after Kammerer's suicide Przibram went
through a crisis. He had lost his oldest and most trusted col-
laborator, and the reputation of his Institute had been gravely
damaged. For several months Przibram seems to have been un-
able to get over the shock. In that confused state of mind he
wrote, in March, 1927, a letter to Nature,
1
in which he attemp-
ted to summarise the whole affair. He listed five detailed 'proofs'
for the genuineness of Kammerer's results; but also seemed to
imply, quoting Bateson's allegations, that concerning events
16
4
from 192,2 onward Kammerer was mistaken in defending the"
'untoward' position of the pad on the doctored specimen:
A picture was taken in September 1922, not in the Biolo-
gische Versuchsanstalt, but in the photographic studio
Reiffenstein, of the well-known specimen, and only from
thence onwards do the mis-statements begin. On the other
hand, up to 1919 the descriptions and figures of nuptial
pads in Alytes given by Kammerer do not fit in with this
specimen ....
We have been able to collect five proofs that in his origin-
al papers Kammerer was not hampered by the doctored
specimen which has invalidated his remarks on the same
subject in his books Inheritance of Acquired Characteristics
(1924) and Neuvererbung (1925). The proofs are as follows:
(1) In Kammerer's original papers the nuptial pad in
Alytes is described and pictured as being 'on the dorsal
side of the thumb and on the thumb-ball' (1909, p. 516, fig.
26a), 'on the dorsal and radial side ofthe first fingers' (1919,
p. 336), and 'across the thumb-ball on the whole internal
side of the fore-arm to near the elbow' (p. 337, tb. x, fig. 2),
in accord with the general appearance of nuptial pads.
Even in 192,3, when Kammerer showed a lantern slide of
the critical specimen before the Zoological Society of
London, he did not mention the disposition of the nuptial
pad on the whole palm of the hand (see Bateson, Nature,
Dec. 22, 1923, and letter to Przibram). It was not until the
photographs of this specimen were used in his books (1924,
p. 53. fig. 9 to the right; 1925, fig. 9. facing p. 20) that
Kammerer mentions and defends the untoward position of
the pad in the palm and on the outer border of the last
(fourth) finger .•..
What these rather obscure passages seem to mean is that the
specimen was already doctored when, in 192,2, the Reiffenstein
photograph was taken, because that photograph shows the
black patch on the palm, which does not fit 'the descriptions
and figures of nuptial pads' as given in Kammerer's earlier
papers, up to and including the last one of 1919- Up to 1919
Kammerer 'was not hampered by the doctored specimen' j
whereas from 1922 onward 'the mis-statements [obviously
Kammerer's mis-statements] begin'; and only after including
16
5
the Reiffcnstein photograph of the doctored specimen in his
books (1924 and 1925) does he 'mention and defend the un"
toward position of the pad in the palm' which up to that date
he had passed over in silence. The inference is that although
the doctoring was done by somebody else, Kammerer was
either unaware of it, or preferred not to mention it. (How
not mentioning the conspicuous broad black mark across the
palm of the specimen on display could help matters neither
Przibram nor Bateson explained.)
The central issue for Przibram is the Reiffenstein photograph,
and on this his letter directly contradicts what he said about it
earlier on. It will be remembered that in August, 1926, he
wrote a parallel report to Noble's published in Nature. In that
report Przibram cited the photograph in question as one of the
main proofs for the presence of the pads in the original state of
the specimen before it had been tampered with:
Fortunately, there are photographic plates in existence
showing the state of the specimen before it left Vienna for
Cambridge and during its stay in England. One of these
photographs was taken in the presence of Dr. J. H.
in the atelier of Reiffenstein (Vienna), and the negative
travelled with Dr. Quastel to England and has been in the
possession of Mr. M. Perkins (Trinity College, Cambridge)
since April 1923. A reprint of it is given in Kammerer's
Neuvererbung, W. Seifert-Verlag,
I925 (Abb. 9, facing p. 20).
He then goes on to quote at length testimonies from Dr.
Quastel, Michael Perkins and W. Farren (a photographic expert
in Cambridge) to the effect that the photograph 'shows no
traces of any manipulation or retouching of the actual image of
Alytes'. He also quotes Kammerer to the effect that 'he remem-
bers the black substance to have been in the same place and
amount, even in the living animal'. In other words, Przibram
accepts that the black mark on the palm, as shown in the photo-
graph, was genuine and that the forgery was done after the
specimen's return from Cambridge.
Thus the same photograph showing the blackened palm is
invoked in Przibram's 19261etter to Nature as evidence for the
defence, and in his 1927 letter as evidence for the prosecution.
What may have caused confusion? No new evidence
had come to light since Kammerer)s death. Przibram must have
I66
seen, countless times, the specimen with its blackened palm and
the Reiffenstein photograph of it, yet apparently it had never
before occurred to him that the pad was 'in the wrong place'.
After Kammerer's suicide, however, and all the nasty rumours
it provoked, his mind must have been milling round and round
the events of the past until they became hopelessly muddled.
He probably went back to the controversy of 1923, saw it in a
different light, and only now discovered that 'the pad was in the
wrong No doubt he had glanced at the letters in Nature
at the time, but he must have dismissed them as irrelevant, just
as he had throughout all these years found nothing wrong with
the black-handed specimen and its photograph. In this state of
mind, Bateson's letter with its allegation that even in his English
lectures Kammerer 'did not mention' the unspeakable location
of the pad, must have seemed to Przibram to clinch the argu-
ment by making Kammerer appear to have used questionable
methods in his later years. Bateson was dead, but his letter
seemed to have had the effect of a minor time-bomb on Przi-
bram) s mind, getting him into a tangle of contradictions.
Even so, he never for a moment suspected that K.ammerer
himself committed the forgery. It is greatly to Przibram's credit
that he felt no resentment toward Kammerer for the damage his
suicide had caused to the Institute, and that even in his dis-
turbed state of mind he concluded the letter with his five
'proofs' of the soundness of Kammerer's earlier results (Le.
(1) that the 1919 basic paper showed no pads in the wrong
places; (2) an earlier photograph in the same paper; (3) the
microtome sections showing the difference between normal and
and (4) showing the difference between Alytes'
pads and pads in other specimens; (5) lastly, K.andler's sections
of the rudimentary pad found in a normal specimen of Alytes).
Przibram was fallible, but of an almost masochistic honesty;
it would have been far better for the prestige of his Institute if
he had simply said that an unknown person had tampered with
the specimens, and left it at that.
There is one more point to be dealt with in this context.
Though Kammerer did mention the blackening of Calmost the
whole hand'at his English lectures, why did he not mention it
in his 1919 paper, since, in his own words, the blackening was
in the same place and amount in the living animal? I think the
answer is implicitly contained in the following quote from the
1919 paper (Kammerer's italics):
The first specimens of Alytes males on heat that I found
equipped with nuptial pads displayed these in the form of
sharply defined, greyish-black thickenings of the upper and
radial sides of the first finger. The second finger does not
show any traces of this formation either macroscopically or
under the magnifying glass. However, the more Alytes
males, particularly those of the F 4 and F 5 generation, came
on heat, the more frequently one could observe that the pad
did not always appear in the original regions; but that,
whether in the same individual or among different indi-
viduals, it has a fairly wide region at its disposal, ranging
from over the thumb-ball and the whole inner side of the
forearm to the proximity of the elbow, and that within this
region the pads show great variability in their extension
and pattern. Even asymmetries do occur: for instance, on
the left only a patch of the forearm might show the pad.
Figure 2, Table X shows a male of the F5 generation at the
height of the mating season displaying a mighty pad over a
large area of the radial side of the forearm and, incidentally,
also over the thumb-balls, leaving, however, the phalanges
unaffected.
Within the same individual the pad area is not irrevoc-
ably fixed to the same spot either; and we can even follow
the direction of the variability: generally speaking, the area
increases from one mating season to the next. If, for in-
stance, the first pad appeared on the finger tip only, then
during the second mating season the whole finger is affec-
ted, during the third the ball of the third, the fourth an
adjacent area of the forearm. In other words, the varia-
bility moves in the direction of increasing areas and a ~ the
same time in the direction from distal towards proXimal
regions.
2
He did not explicitly mention that in one (or several) speci-
mens the mark had also spread across the palm; in view of the
extreme variability of the pads both in Alytes and in other
species, he probably thought this superfluous, as the passage
quoted implies such a possibility. Since some toads develop
pads on the tips of their fingers, others on their hind-legs, to
talk of 'the wrong place' seems hardly defensible, and the whole
controversy looks like making a mountain out of a mole hill. But
it had to be included in this account for the sake of completeness.
r68
APPENDIX 5
Cion a
The Ciona experiments have been briefly summarised on
PP·4S
f
.
In his I923 Cambridge and Linnean lectures Kammerer said:
... I carried out, before I9I4, what may really be an ex peri-
mentum crucis. I have written a few words on it in my
Allgemeine Biologie. There has been no detailed publication
as yet. The subject is the Ascidian, Ciona intestinalis. If one
cuts off the two siphons (inhalant and exhalant tubes), they
grow again and become somewhat larger than they were
previously. Repeated amputations on each individual speci-
men give finally very long tubes in which the successive
new growths produce a jointed appearance of the siphons.
The offspring of these individuals have also siphons longer
than usual, but the jointed appearance has now been
smoothed out. When the nodes are to be observed, they are
due not to the operation but to interruptions in the period
of growth, just as in the winter formation of rings in trees.
That is to say, the particular character of the regeneration
is not transferred to the progeny, but a locally increased
intensity of growth is transferred. In unretouched photo-
graphs of two young Ciona attached by their stolons to the
scratched glass of an aquarium, the upper specimen is
clearly seen to be contracted; the lower is at rest artd shows
its monstrously long siphons in full extension. They were
already there at birth, for it was bred from parents the
16
9
siphons of which had become elongated by repeated ampu-
tation and growth.
1
On November 3 Nature published a letter from H. Munro
Fox of the Cambridge Zoological Department, of which the
relevant passages read:
I repeated these amputation experiments between June
and September last at the Roscoff Biological Station. The
oral siphon was removed from 102 Ciona intestinalis which
were growing attached to the walls of the tanks. The ani-
mals varied in length from 0'9 to 4'8 cm. As controls, 235
unoperated individuals were kept under observation. In
none of the operated animals was there any further growth
of the siphons after the original length had been re-
attained .... In 1913 it was shown at Naples that ab-
normally long siphons of Ciona intestinalis can be grown
by keeping the animals in suspensions of abundant food
(Bioi. Centrbl. 1914. vol. 34, p. 429). Were this the reason
for the long siphons of Dr. Kammerer's operated Ciona,
it should have been clear from controls of unoperated
animals kept in the same water.
MacBride replied to Munro Fox's letter on November 24 in
Nature (his italics):
As Dr. Kammerer took a deep interest in the projected
repetition of his experiments on Ciona, and wrote to me
twice this summer to learn if repetition were being attemp-
ted and under what conditions, perhaps you will allow me
to make some remarks on Mr. Fox's letter, as Dr. Kam-
merer is now in America.
Dr. Kammerer, whilst in Cambridge, wrote out a full
account of the precautions to be observed in making these
experiments. At that time he did not know that Mr. Fox
was going to take up the work: another Cambridge biolo-
gist had undertaken to do so, but this gentleman was pre-
vented by illness from doing the work. To him, however,
Dr. Kammerer had transmitted his information. I under-
stand-Mr. Fox will correct me if I am wrong-that Dr.
Kammerer's instructions did not reach Mr. Fox. In these
circumstances it is not surprising to learn that Mr. Fox
failed to obtain Dr. Kammerer's results, since he has
170
L
.'
tumbled into one of the most obvious pitfalls. It may sur-
prise him very much to learn that Dr. Kammerer got the
same results as he did when, like Mr. Fox, he cut off only the
oral siphon. Since the anal siphon remains of normal
length and the reaction is of the animal as a whole, the re-
generated oral siphon is of normal length also. But when
both anal and oral siphons are amputated in a very young
animal, then long siphons are regenerated. I have a photo-
graph which shows an operated Ciona and a normal one
growing side by side in the same tank, and the contrast
between the lengths of their siphons is obvious. When Dr.
Kammerer returns from America I hope that Mr. Fox will
communicate with him and repeat the experiments, ob-
serving Dr. Kammerer's precautions, when, I feel con-
fident, he will obtain Kammerer's results.
Kammerer, on his return from America, also replied to
Munro Fox-Nature, December 8 (his italics):
In Nature of November 3, page 653, Mr. H. Munro Fox
announces that he did not succeed in repeating my results
in his Ciona experiments in Roscoff: amputated siphons
regained only their normal length. Mr. Fox supposes that
the extra growth in length of the siphons in my experiments
was produced by extravagant feeding, and not by the re-
generative activities of the animals.
Before Mr. Fox publishes the full account of his work,
which he promises, I beg him to note tlle following facts,
namely:
(I) The two principal cultures (operated and control) of
my CiQna were placed at the same time and at the same
stage of development, with the same provision of food, in
two precisely similar aquaria, which stood beside each
other. The dimensions of these aquaria were 300 x 170 x
100 centimetres. I did not undertake a quantitative estima-
tion of the number of micro-organisms present; but the
food available was, so far as I could see, rather on the
scanty than on the abundant side.
All the specimens in the control culture possessed short
siphons, and therefore the influence of food on the length
of siphon is excluded.
(2) I am not the first and only observer who has noted
the 'super-regeneration' of the siphons after they have been
17
1
cut off several times. Mingazzini* asserts that siphons
amputated three or four times at intervals of a month be-
came longer after each regeneration. Mingazzini was able
in this way to produce artificially the local variety, 'macro-
siphonica', found in the Gulf of Naples. I fully anticipated
that the decisive experiment on regeneration and inherit-
ance in Ciona would encounter violent contradiction. On
that account I took care to construct this critical experiment
out of experiments which had already been made by other
investigators. That this was possible in the case of Ciona
was one of the reasons which led me to choose this species.
Indeed, I have had a predecessor (E. Schulz) also on the
question of the regeneration of the 'Keimplasma' out of
somatic material, though his experiments were made not
on Ciona but on another Ascidian (Clavellina). The only
originality which I claim is the combination of well-known
experiments and their application to the solution of a prob-
lem of inheritance.
Barfurth, t after he had discovered (at that time in his
laboratory at Dorpat) that the limbs of frog-larvae had the
power of regeneration, laid stress on the superiority of one
positive result as against any number of negative results.
'Even if only Dorpat tadpoles regenerated their limbs,
nevertheless his result would be established.' I make the
same claim for Ciona, 'even if only C£ona from Naples and
Trieste grow long siphons'. Finally, have perhaps only
southern populations this power?
On December 22 of the same year, Przibram wrote to Bateson
refusing the '£25 in the same letter he also mentioned
Ciolla:
2
In ease you have noticed Mr. Munro Fox's letter in Nature,
No. 2818, on Ciona, I would like to direct your attention to
the fact that the discovery of its siphons lengthening with
repeated removal was not made first by Kammerer. It was
known so long ago as I897 by Mingazzini's experiments,
which were, in their turn, based on a previous observation
"" 'Sulla regenerazione nei Tunicata', Bolletino Soc. Nat. Napoli, Ser. I,
year 5. 189I. (An abstract of this paper appeared in the Naples
Zoologisclzer Jah"esberieht for 1891 under the heading 'Tunicata'.)
t 'Sind die Extremitliten der regenerationsfahig?' Arch Entw-
Meeh., vol. I, 1894.
r
of our friend in common, Jacques Loeb, as he mentioned to
me in I907 during my stay in California. So I do not sec how
Mr. Fox's inability to reproduce the experiment allows him
to deny Kammerer's success with the first generation.
On January 5. 1924, B. Stewart, a student at Trinity and
member of the Cambridge Natural History Society, also wrote
about Ciona to Nature. He was an amateur photographer,
who had taken pictures of Kammerer's specimens and copied
Kammerer's photographs of Ciona.
There are three photographs of Ciona. The first is of a
single untreated specimen, the second of a group showing
artificially produced var. macrosiphonica, and the third of
two untreated offspring of the latter. In view of the various
magnifications, both in the camera and from perspective,
and since the whole of the animal is not visible in most
cases, simple measurements would be meaningless. How-
ever, the increase of the siphon of v. macrosiphonica is
chiefly in the direction of length, and therefore the ratios of
length to breadth of the siphons provide a satisfactory
method of comparing the specimens. The ratios are:
Photograph 1. (Untreated, fully extended specimen.)
Oral Siphon I'9, aboral 1·65.
Photograph II. (Group.)' In a single fully extended
specimen, doubtless that referred to by Prof. MacBride,
the ratios are 2'0 oral and 1'65 aboral. In the remainder the
ratios when expanded are 4'0 to 4'3 oral and 2'0 to 4'3
aboral, and when contracted 2'4 oral and 1'9 aboral.
Photograph III. One of these two young offspring of
v. macrosiphonica is completely expanded or nearly so, the
other is quite contracted; in the former the ratios are 4' I
oral and 2'05 aboral, in the latter they are 2'35 and 1'4,
The validity of the means qf comparison suggested
above is shown by the ratios of length to breadth for the
main part of the body lying, in all the four or five speci-
mens in which it can be measured, between 4'1 and 4.8;
i.e. the error due to varying expansion, position, and focus
cannot possibly be more than 20 per cent, yet v, macro-
siphonica shows an increase in length of the siphons of as
much as 125 per cent.
On January 19 J. T, Cunningham of East London College
173
took issue with MacBride's explanation of the reasons for
Munro Fox's negative results. He looked up Mingazzini's paper
on the subject and found that 'it is distinctly stated that in some
cases the buccal and cloacal siphons were cut off in different
individuals, sometimes in the same individual, and that in
either method a regenerated siphon showed increased length.
It is to be noted that Dr. Kammerer in his letter in Nature of
December 8 does not confirm the statement of Prof. MacBride
in the issue of November 24.'
MacBride replied to this-Nature, February 9, 1924.
In Nature of January 19, p. 84, there appears a letter from
Mr. Cunningham in reference to the regeneration of the
siphons of Cion a, in which he calls in question a statement
of mine in a letter in the issue of November 24. In my
letter I attributed the failure of Mr. Fox to get lengthened
siphons after amputation to the fact that he cut off only the
oral siphon.
Mr. Cunningham says that Dr. Kammerer did not con-
firm my view in his subsequent letter to Nature (which
incidentally I translated for him and sent to Nature). This
is true; but I received afterwards a letter from Dr. Kam-
merer in which he explicitly agrees with my explanation
and says that he had not realised that Mr. Fox had only cut
off one siphon.
It appears that Mingazzini-about whose work Mr.
Cunningham learnt from the letter which I translated-
succeeded even when he cut off only one siphon. It may,
therefore, be the case, as Dr. Kammerer suggested, that
Mr. Fox failed, not because he cut off only one siphon, but
because he was dealing with a northern race of Ciona.
The importance of the reference to Mingazzini's work
lies in this, that this work unequivocally supports Dr.
Kammerer's statements: many were inclined to doubt their
trustworthiness after the publication of Mr. Fox's letter.
And there the controversy came to rest-as the others did.
Whatever the reason for Munro Fox's failure to obtain elon-
gated siphons, his negative result has to be weighed against the
positive results obtained by Mingazzini, Jacques Loeb and
shown on Kammerer's photographs. Nothing that transpired in
the controversy justifies the abandonment of a line of research
with far-reaching theoretical implications.
174
I
I'
t
REFERENCES
CHAPTER ONE (pages 13 to 26)
I. Neue Freie Presse, Vienna, September 25. 1926. 2. October 30,
1926, p. 635 f. 3· K. Przibram (1959), p. 185. 4. Kammerer (1899
and 1900). 5. H. Przibram (1926), pp. 401 if. 6. Private communica-
tion, April 6, 1970. 7. Goldschmidt (1949), p. 22. 8. The Bateson
Papers. 9. MacBride (1924), p. 88.10. Kammerer (1914b), pp. la-II.
II. Cannon (1959), p. 45. 12. Kammerer (1906, 1909a, 1919a).
CHAPTER TWO (pages 27 to 38)
I. Goldschmidt (1949), p. 221. 2. Kammerer (I914b), pp. 15-16.
3· Hardy (1965), p. 156. 4· Waddington (1952). 5. Bergson (I9II),
pp. 44-5, quoted by Himmelfarb (1959), p. 369. 6. Cannon (1959).
pp. ix-x. 7. Butler (1951 ed.), p. 167. quoted by Himmelfarb (1959).
p. 362. 8. Butler (1879). p. 54. 9. Darlington in preface to reprint of
On the Origin of Species (1950). 10. Life and Letters, II, p. 215.
II. Quoted by Himmelfarb (1959). 12. Darwin (1868). 13. Third
Letter to Bentley, Opera Omnia, IV, p. 380. 14. Quoted by Hardy
(1965). p. 157. 15· Hardy, op. cit., p. 159. 16. Darlington (r953),
pp. 219-21 •
CHAPTER THREE (pages 39 to 47)
I. Kammerer(1904),PP. 165-264. 2. Kammerer (1907c), pp. 99-102.
3· Kammerer (Ig07b), p. 34· 4· Kammerer (1923), p. 637· 5.
Kammerer (1925), p. 45. 6. Ibid., p. 60. 7. Ibid., pp. 48-9. 8.
Kammerer (1923a), p. 639. 9. Bateson, letter in Nature, May 16,
1923. 10. Bateson (1913), p. 201. II. MacBride, letter in Nature.
January 17, 1925. 12. Kammerer (1919a), p. 327. 13. MacBride,
letter in Nature, December 5, 1925. 14. Goldschmidt (1940), p.
2057 n. IS. Goldschmidt (1949), p. 221.
CHAPTER FOUR (pages 48 to 58)
I. The Bateson Papers, July 17, 1910. 2. H. Przibram (1926).
3. Bateson (1928). 4· Private communication, April 8, 1970.
S. Bateson (1902). 6. Hardy (1965), p. 89.
175
CHAPTER FIVE (pages 59 to 64)
I. Bateson (1913), p. I9I. 2. Ibid., p. 190. 3. Ibid., p. 227· 4. Ibid.,
p. 199. 5. Ibid., p. 202. 6. The Bateson Papers.
CHAPTER SIX (pages 65 to 72)
1. Kammerer (I919a), pp. 357 ff. 2. Kammerer (1925), p. 34·
3. Darlington (1953). p. 222. 4. The Bateson Papers. 5. The Bateson
Papers. 6. Ibid. 7. Ibid.
CHAPTER SEVEN (pages 73 to 86)
I. MacBride, letter in Nature, June 23, 1923. 2. Kammerer (1919a),
p. 328. 3. Kammerer (I923a), p. 640. 4· The Bateson Papers.
5. Montagu (1970), p. 139.6. Private communication, April 6, 1970 •
7. June 23, 1923. 8. Letter in Nature, August 18, 1923. 9. The
Bateson Papers. 10. Kammerer, letter in Nature, August 18, 1923.
II. Letter in Nature, September 15, 1923. 12. Letter in Nature,
August 18, 1923. 13. Letter in Nature, December 8, 1923.
CHAPTER EIGHT (pages 87 to 97)
I. The Bateson Papers, September 20, 1923. 2. Minutes of the
Council of the Cambridge Natural History Society, April 27, 1923.
3. Ibid. 4. Ibid. 5. Ibid., May I, 1923. 6. The Bateson Papers,
October I, 1923. 7. Ibid., February 27, 1924. 8. Lecture announce-
ment, F'ebruary 16, 1924, New School for Social Research, New
York. 9. Kammerer (1926b). 10. July 16, 1927.
CHAPTER NINE (pages 98 to II6)
I. Letter in Nature, October 16, 1926. 2. Nature, August 7, 1926.
3. H. Przibram, letter in Nature, April 30, 1927. 4. Lataste (1876).
5. The Bateson Papers, August 14, 1920. 6. April 30, 1927. 7. Letter
in Nature, August 21, 1926. 8. April 30, 1927. 9. Nature, August 17,
1926. 10. H. Przibram (1926). II. Letter in Nature, April 30, 1927.
12. K. Przibram (1959), p. 188. 13. Private communication, October
19,197°. 14· Ibid., October 27,1970. IS. Neue Freie Presse, Septem-
ber 27, 1926.
CHAPTER TEN (pages II7 to 122)
I. R. Wettstein, Neue Freie Presse, December 16, 1926. 2. Ibid.
3. Ibid. 4. September 25, 1926.5. Der Abend, September 24, 1926.
6. Private communication, July 2, 1970. 7. K. Przibram, op. cit.,
P·
18
7·
EPILOGUE (pages 123 to 134)
1. Kammerer (1923a), p. 639. 2. Bateson (1924), p. 405. 3. Ibid.
p. 406. 4. Bateson (1913), p. 248. 5· Johannsen (1923), p. 140.
6. June 26, 1970. 6a Thorpe (1969), p. 1. 7. von Bertalanffy (1969),
pp. 66-7. 8. Koestler (1967), pp. 158-1).9. Waddington (1957), p. 182.
10. Ibid. 1 I. New York Evening Post, February 23, 1924.
176
~
I
I
APPENDIX I (pages 135 to 143)
I. Jung (196o), p. 420.2. Ibid., p. 438.3. Kammerer (1919b), p. 24.
4. Ibid., p. 25· ¥. Ibid., p. 27. 5. Ibid., p. 36. 6. H. Przibram (1926).
7· Kammerer (1919b), p. 93. 8. Ibid., p. 137.9. Ibid., p. 165. 10. Ibid.
p. 454· II. Ibid., p. 456. 12. Jung, op. cit., p. 441. 13. Ibid., p. 435.
14. Kammerer (1926a).
APPENDIX 2 (pages 144 to 147)
1. Goldschmidt (1949), pp. 220-2. 2. Cannon (1959), p. 46. 3. Ibid.
4. Goldschmidt (1949), p. 221.
APPENDIX 3 (pages 148 to 159)
1. The Bateson Papers. 2. Ibid., 3. Ibid. 4. Bateson (1913), pp. 207-8.
5. E. G. Boulenger (19II), p. 323. 6. G. A. Boulenger (1912), pp.
572-3. 7. Ibid., p. 579. 8. Ibid. 9. The Bateson Papers. 10. G. A.
Boulenger (1917), pp. 173-4. II. G. A. Boulenger (1912). 12. G. A.
Boulenger (1917), pp. 174--5. 13. Ibid., pp. 177-8. 14. Ibid., p. 176.
IS· Letter in Nature, July 3, 1919. 16. G. A. Boulenger (1917). p. I77.
17. Ibid., p. 180 f. 18. Kammerer (1914a), p. 260.19. G. A. Boulenger
(1917), p. 181. 20. Kammerer (1906), pp. 68-9. 21. Kammerer
(1909a), pp. 475-6.
APPENDIX 4 (pages 160 to 168)
I. April 30, 1927. 2. Kammerer (I919a), pp. 336-7.
APPENDIX 5 (pages 169 to 174)
I. Kammerer (1923a), p. 369. 2. LetterinNature, December 22,1923.
177
/,
~
II
BIBLIOGRAPHY
1. WORKS BY PAUL KAMMERER*
(Not including popular articles, lectures and pamphlets on socio-
cultural subjects.)
Technical Papers
Die Reptilien und Amphibien der hohen Tatra. Mitteilungen der
Sektion fUr Naturkunde des bsterreichischen Touristenklub, XI,
Heft 6 und 7, 1899.
HaJtzeher in GeJangenschaJt. Natur and Haus, VIII, Hefte 22 und 23,
19°0.
Beitrag zur Erkenntnis der VerwandtschaJtsverhiiltnisse von Salaman-
dra atra und maculosa. Arch. 1904. 17. 165-264.
Vber die Abhiingigkeit des Regenerationsvermogens der Amphibien-
larven von Alter, Entwicklungsstadium und speziji.scher Grosse.
Arch. 1905. 19. 148-80.
Experimentelle Veriinderung der Fortpjlanzungstiitigkeit bei Geburts-
helferkrote (Alytes obstetricans) und Laubfrosch' (Hyla arborea).
Arch. 1906a. 22. 48-14°.
Kunstlicher Melanismus bei Et"dechsen. Zbl. f. Physio!., Leipzig, 1906b.
20. 261-3.
Bastardierung von Flussbarsch (Perca jluviatilis L.) und Kaulbarsch
(Acerina cernUa L.) Arch. 1907a. 23. 5I1-51.
Vererbung erzwungener Fortpjlanzungsanpassungen. I u. II. Mitteil-
ung: Die Nachkommen der spatgeborenen Salamandra maculosa und
der Jruhgeborenen Salamandra atra. Arch. 1907b. 25· 7-51.
Vererbung der erworbenen EigenschaJt habituellen Spiitgebiirens bei
Salamandramaculosa. Zbl. f. Physiol., Leipzig, 1907c. 21. 99-102.
Erzwungene FortpJlanzungsveriinderungen und deren Vererbung, etc.
Zbl. f. Physiol., Leipzig, 1907d. 21. Nr. 8.
Symbiose zwischen Libellenlarve und Fadenalge. Arch. 1908a. 25.
52- 81.
Regeneration sekundiirer Sexualcharaktere bei den Amphibien. Arch.
1908b. 25. 82-124.
:II< Arch. refers to: Archiv fur Entwicklungsmechf.Jnik der Organismen,
Leipzig.
179
Regeneration des f)ipterellfiiigels beim Imago. Arch. 1908c. 25. 349-60.
erzwlIlI/.:cller Fort pfiallzungsanpassungen. III. 1l1itteilung:
Die Nachlw1I1mcn der nicht brutpfiegellden Alytes obstetricans.
Arch. 19o(ja. 28. 41-7-5+6.
Allgemeillc Symhiose lind Kall/pf ums Dasein als gleichberechtige Trieb-
Krafte der E'volutio1/. Arch f. Rssen- u. Gesellsch.-Bio!., Leipzig
und Berlin, 1909b. 6. 585-608.
Vererbung erZWUll.r;ener Farb- Imd Fortpjlanzzwgsveriinderungen.
Natur, Leipzig, 1909c. I. 94-7.
VererbunJ.? erzwun,c:ellcr Farbveranderungen. I. u. II. Alitteilung,'
Induktion von weiblichem Di11lorphismus bei Lacerta muralis, von
miinnlichem Dimorphismus bei Lacerta fiumana. Arch. I91oa. 29.
45
6
-98.
Die Wirkung iiusserer [,ebensbedingzmgen auf die organisdze Variation
im [,ichte der e.\j}erimentellen Jlilorphologie. Arch. 1910b. 30. 1.
379-4
08
.
Das Beibehalten jugendlich unrezfer Formzustiinde (Neotonie und
ProJ.?enese). Ergcbn. d. wisscnsch. Med., Leipzig, 1910C. 4. 1-26.
Gregor Mendel lind seine Vererbungslehre mit Rucksicllt aUf ihre
Bedeutung fiir die medizinische Wissenschaft. Wien. med. W chnschr.
1910d. 9. 23 67-72 .
Beweise Jiir die Vererbullg erwurbeller Eigenschaften durch planmii.ssige
Zuchtung. Deutsche Gescllschaft fur Zuchtungkunde, Berlin,
I9Ioe.
Vererbung kunstlicher Zeugungs- und Farbveriillderungen bei Reptilien.
Vortrag Internat. Physio!. Kongress, Wien. Umschau. XV. Nr. 7.
133-56. I9Ila.
Mendelsche Regeln und Vererbung erworbener Eigenschaften. Verhandl.
d. Naturforsch. Ver. Brunn. 191 lb. XLIX (Mendel-Festband).
Experimente iiber Fortpfianzung, Farbe, Augen und Korperreduktt"on
bei Proteus anguineus Laur. (zugleich: Vererbung erzwungener
Farbveriinderungen. III. Mitteilung). Arch. I9I1- 12. 33· 349-461.
Di1'ekt induzierte Farbanpassungen und deren Vererbung. Zeitschrift
indukt. Abst.-u. Vererbungs!. 191 I. IV. 279-88 und Verh. VIII
Internat. Graz. 1912.263-71.
Experimente uber Fortpfianzung, etc. IV. Mitteilung: Das Farbkleid
des Feuersalamanders, Salamandra maculosa Laurenti in seiner
Abhiingigkeit von der Umwelt. Arch. I913a. 36. 4-193.
Nachweis normaler Funktion beim herangewachsenen Lichtauge des
Proteus. Arch. f. d. ges. Physiol., Bonn, 19I3b. 51. 1090-4.
Bemerkungen zum Laichgeschiift und der Brutpfiege bei der Geburts-
helferkrote (Alytes obstetricans). Blatter fUr Aquarien u. Terrari-
enkunder. 1914a. XXV. Nr. IS. 259-61.
Die Bedeutung der Vererbung erworbener Eigenschaften fur Erziehung
und Unterricht. Wien, I9I4b.
Vererbung erzwungener Formveriinderungen. I. Mitteilung: Brunst·
180
II
I
i
[
I
>
j
I
C
I
schwiele der Alytes-Mannchen aus 'Wassereiern' (Zugleich: Verer-
bung erzwungener Fortpjiallzungsanpassungen. V. Mitteilung). Arch.
19
I
9
a
. 45· 32 3-70 .
Die Zeichnung von Salama1ldra maculosa in durchfallendem farbigem
Licht. Arch. 1922. 50. 79-107.
Zuchtvel'suche iiber Vererbung erworbener Eigenschaften. Natur,
Leipzig. 1922-3. 14. 305-I1.
Breeding experiments on the inheritance of acquired characters. Nature.
1923a. III. 637-40.
Methoden der experiment ellen Variationsforschung. Hand. d. bioI.
Arbeitsmeth., Berlin, 1923b. Abt. 9 T.3.
Das Darwinmuseum zu Moskau. Monistische Monatshefte, October,
1926a. 1 I. 377-82.
Methoden und Zuchtung von Reptilien und AmphibietZ. Pjiege und Zucht
weiterer wirbelloser Landtiere. In: Handb. d. bioI. Arbeitsmeth.,
Berlin, 1928. Abt. 9 T. I, 2, I.
Books
Bestimmung und Vererbung des Geschlechtes bet' Pjlanze, Tier und
Mensch. Leipzig, 1913.
Genossenschajten von Lebewesen auf Grund gegenseitiger Vorteile
(Symbiose). Stuttgart, 1913c.
Allgemeine Biologie. Stuttgart, 1915 (3. Aufl.. 1925).
Das Gesetz der Serie. Stuttgart, 1919b (2. Aufl.. 1921).
The Inheritance of Acquired Characteristics. New York, 1924.
N euvererbung oder Vererbung erworbener Eigenschaften. Stuttgart, 1925.
Der Artenwandel aUf Inseln und seine Ursachen. Leipzig und Wien,
1926b.
2. THE KAMMERER CONTROVERSY IN N ature*
1919 May 22: MacBride (L)
July 3: Bateson (L)
1923 May 12: Kammerer (A)
May 26: Cunningham (L)
June 2: Bateson (L)
June 23: MacBride (L)
June 30: Bateson (L)
July 21: MacBride (L)
July 28: Cunningham (L)
August 18: Kammerer (L)
Perkins (L)
September 8: MacBride (L)
Sir Arthur Keith (L)
September 15: Bateson (L)
November 3: Munro Fox (L)
... A refers to articles, L to letters.
181
November 24: MacBride (L)
December 8: Kammerer (L)
December 15: Cunningham (L
December 22: Przibram (L)
1924 January 5: Stewart (L)
January 19: Cunningham (L)
February 9. MacBride (L)
May 17: Dover (L)
June 5: CaIman (L)
July 26: Cunningham (L)
September 6: Przibram (L)
September 27: CaIman (L)
1925 January 10: MacBride (L)
November 28: MacBride (L)
December 5: MacBride (L)
1926 January 9: Cunningham (L)
February 13: MacBride (L)
March 6: MacBride (L)
March 20: Cunningham (L)
May 29: Cunningham (L)
August 7: Noble (A)
Przibram (A)
August 21": MacBride (L)
October 2: Obituary notice
October 9: Noble (L)
October 16: Obituary by Przibram
October 30: Obituary (unsigned)
November 6: MacBride (L)
1927 April 30: Przibram (L)
Kiplinger (L)
May 14: MacBride (L)
July 16: MacBride (A)
3. OTHER WORKS MENTIONED IN THIS BOOK
Bateson, W., Mendel's Principles of Heredity: A Defence. Cambridge,
19°2.
--Problems of Genetics. New Haven and London, 19
1
3.
--Naturalist. Cambridge, 1928.
--" Letters from the Steppe. London, 19z8.
The Bateson Papers, Library of the American Philosophical Society,
Philadelphia.
Bergson, H., Creative Evolution. Tr. A. Mitchell. London, 19
11
•
von Bertalanffy, L., in Beyond Reductionism. See Koestler and
Smythies, ed., 1969.
Boulenger, E. G., Proc. Zool. Soc., 1911, p. 3
2
3.
182
11
:!
Boulenger, G. A., 'Observations sur l'accouplement et la ponte de
l'AIyte accoucheur, Alytes obstetricans'. Academic Royale de
Belgique, Bulletin de la Classe des Sciences, 1912. Nos. 9-10.
570 -9.
--'Remarks on the Midwife Toad (Alytes obstetricans), with
reference to Dr. P. Kammerer's Publications.' Annals and
Magazine of Natural History, August, 1917. Ser. 8, Vol. XX.
173-84.
Butler, Samuel, Evolution Old and New. 1879.
--Notebooks. Ed. G. Keynes and B. Hill. New York, 1951.
Cannon, H. Graham, The Evolution of Living Things. Manchester,
1958.
--Lamarck and Modern Genetics. Manchester, 1959.
Darlington, C. D., in preface to On the Origin of Species. Reprint of
1st ed., London, 1950.
--The Facts of Life. London, 1953.
Darwin, Charles, The Variation of Animals and Plants under
Domestication. 2 vols. London, 1868.
--On the Origin of Species. Reprint of 1st ed., London, 1950.
Dobzhansky, T., The Biology of Ultimate Concern. New York, 1967.
Flammarion, c., L'Inconnu et les Problemes Psychiques. Paris, 1900.
Focke, W., Die Pflanzen Mischlinge, 1881.
Goldschmidt, R., The Material Basis of Evolution. New Haven, 1940.
--'Research and Politics'. Science, March 4, 1949.
Hardy, Sir A., The Living Stream. London, 1965.
Himmelfarb, G., Darwin and the Darwinian Revolution. London,
1959·
Johanssen, W., 'Some Remarks about Units in Heredity'. Hereditas,
1923. IV, p. 140.
Jung, C. G., The Structure and Dynamics of the Psyche. The Collected
Works, Vol. 8. Tr. R. F. C. Hull. London, 1960.
Kammerer, P.-see Bibliography Part, I.
Koestler, A., The Sleepwalkers. London and New York, 1959.
--The Act of Creation. London and New York, 1964.
--The Ghost in the Machine. London and New York, 1967.
--ed. with J. R. Smythies, Beyond Reductionism-New Perspec-
tives in the Life Sciences. The Alpbach Symposium. London and
New York, 1969.
Lamarck, J. P., Philosophie Zoologique, 2 vo}s., ed. C. Martins. 2nd
ed., Paris, 1873.
Lataste, F., Ann. Sci. Nat., 1876. (6), tom. 3.
MacBride, E. W., An Introduction to Heredity. London, 1924.
MacDougall, W., British Journal of Psychology. 1927. Vol. 17. 268-
3°4·
Mahler-Werfel, A., MeinLeben. Frankfurt a. M., 1960.
Mendel, G., Experiments in Plant Hybridisation, 1865.
Montagu, 1., The Youngest Son. London, 1970.
Newton, Sir I., Opera Omnia. London, 1779-85.
Noble, G. K., The Biology of Amphibia. New York and London,
1931•
Przibram, H., 'Paul Kammerer als Biologe'. Monistische Monatshefte,
November, 1926, pp. 401-5.
--Experimental-Zoologie. 7 Bande. Vienna and Leipzig: Deuticke,
19
0
7-30 •
Przibram, K., 'Hans Przibram' in Grosse Osterreicher, Band XIII.
Zurich, Leipzig, Vienna, 1959.
St. Hilaire, G., Philosophie Anatomique. Paris, 1818.
Salisbury, F. B., 'Natural Selection and the Complexity of the
Gene', Nature, October 25, 1969. pp. 342-3.
Semon, R., The Mneme. London, 1921.
Smith, J. M., Nature, 1970 • 225. 563.
Smythies, J. R.-see Koestler (1969)'
Sonneborn, T. M., 'Gene Action in Development'. Proc. Royal Soc.
of London B, December, 1970. Vol. 176. No. 1044.
Spetner, L. M., 'Natural Selection versus Gene Uniqueness'.
Nature, June 6, 1970 , pp. 948-9.
Thorpe, W. H., in Beyond Reductionism, see Koestler and Smythies
ed.
Waddington, C. H., The Listener, London, February 13, 1952.
--The Strategy of the Genes. London, 1957.
Whyte, L. L. Internal Factors in Evolution. London, 1966.
•
f
I.
I
INDEX
AGAR, W. R, 36
Altenberg, Peter, 117
BALTIMORE, D., 127
Barfurth, 172
Bateson, Beatrice,s I, 70
Bateson, Gregory, 24, 51, 54,
73, 82, 98, 10" 121
Bateson, W i H i a ~ , 24, 31,41,
44, 45, 49, 50-I, 52, 53-5,
59-
6
4, 65, 66, 67, 68-9, 70-1,
73, 75, 7
6
, 7
8
, 79, 82-6, 87.
9
2
, 93, 94, 96, roo, 101,
102, 103, IDS, 106, 109, 12I,
I25, 126, 147, 148-50, lSI,
15
2
, 153, 154, 156, 157, I59,
160-4, 166, 167, 172
Bauer, Erwin, 50, 100, 157
Beer, Sir Gavin de, 3
2
Bergson, Henri, 31
Bertalanffy, Ludwig von, 74,
12
9, 132
Bles, R J., 77, 148
Barradaile, N., IDS
Boulenger, E. G., 24, 68, 7
1
,
7
2
, 81, 85, 86, 100, 101, 102,
10
3,
10
4. 150, 156-']
Boulenger, G. A., 24, 44, 103,
148-59, 161
Butler, Samuel, 31-2
CANNON, H. Graham, 26, 31 ,
105, I46-']
Chardin. Teilhard de, 129
Congdon, E. B., 123
Carrens, K. E., 53
Crick, Francis, 127
Cunningham, J. T., 83, 173-4
DARLINGTON, C. D., 32,
36-8, 67, 146
Darwin, Charles, 14. 31, 32-3,
34. 51, 52, 53, 57, 93, 96,
12
5-6, 130, 133
Darwin, Francis, 52
De L'Isle, A., 152, 153
Demours, 151, 152
Dobzhansky, 129
Driesch, Hans, 66
EINSTEIN, Albert, 31, 81, 93,
139, 142
FARREN, W., 166
Finkler, Walter, 22
Fisher, Sir Ronald, 55-6
Fbnnnarion,CaInille,I3S
Fliess, Wilhelm, 141
Focke, Wilhelm, 57
Fontgibu, M. de, 135
Fox, Munro, 46, I70-1, 172-3,
174
Freud, Sigmund, IS, So, 141
Frisch, Karl von, 22
GADOW, H., 75,76,77,81, 104,
162
Galileo, Galilei, 34
18S
Galton, Francis, 32, 52, 54
Gardiner, John Stanley, 75, 91 ,
105
Garstang, W., 130
Goethe, 133. 141
Goldschmidt, Richard, 24, 27,
40, 47, 144, 147
Gutmann, Willy von, 1I8
HALDANE, J. B. S., 77, 105
Hardy, Sir Alister, 30, 36, 52,
56, 130
Hanner, Sir Sidney, 85, 105
Hartmann, 151, 153, 154
Huber, Rudolf, 117
Hutchinson, G. Evelyn, 76-7,
89, 105, 148
Huxley, Sir Julian, 30, 129
Hyden, Holger, 109-12, II3,
123, 124
JAMES, William, 130
Jenkins, Fleeming, 52
Johannsen, W., 126-7
Jung, C. G., 135-6, 139,
142-3
KAMMERER'S brothers, 19
Kammerer, Charley, 19. 50
Kammerer, Felicitas, 48, 49,
73, 94, 95, II7, u8, 137
Kammerer, Karl, 18, 19, 20
Kammerer, Lacerta, IS, 19,
20, 49, 54, 94, 95, 1I7, u8,
138
Kammerer, Sofie, 18, 19, 20
Kandler, R., 103. 123, 167
Kepler, Johannes, 34-
Kermauner, Olga, 101, 123
Kiplinger, Walter C., 107
Knoblauch, 40
Koestler, A., 14, 51, 76, ro8,
129, 131, 132
Komarovska, Countess, 121
Kopany, N., 22
Krech, David, 36
186
LAMARCK, Jean Baptiste, 14,
27, 3
1, 3
2
, 34, 59, 133
Lataste, Fernand, 102, 153, 161
Lebrun, 151
Leydig, 153
Loeb, Jacques, 22,46, 173, 174
Lunacharsky, Anatoly
Vasilievich, 15, 144, 145-6
Lysenko, Trofim, 146
MACBRIDE, E. W., 47, 67-8,
69, 72, 75, 76, 7
8, 81, 83. 85,
87-8, 97, 98, 100, 102, 105-6,
160-1, 170 - 1, 173, 174
MacDougall, William, 35-6
Mahler, Gustav, 20, 21
Mahler-Werfel, Alma, 20-1,
22,94
Matthews, L. Harrison, 80-1,
105
Megeeren, van, 107
Megusar, Franz, II4
Mendel, Gregor, 53.54, 55-7,
126, 130
Mingazzini, N., 46, 172, 174
Monk, J. L., 150
Montagu, the Hon. Ivor, 81-2,
89
Morgan, T. H., 92
Muller, Lorenz, 97
NEWTON, Sir Isaac, 34. 53
Nietzsche, Friedrich Wilhelm,
141
Noble, G. K., 98-9, 100, 101,
102, 103, 104, 105, 106, 107,
108, Il2, II3, II4, II5-
I6,
uS, 121, 124, 147, 166
Nuttall, G. H. F., 91, IDS
ONSLOW, the Hon. Mrs., 75-6,
105
PAULI, Wolfgang, 142
Pavlov, Ivan Petrovich, 17,
34-5, II6
Pearson, Karl, 54, 55, 56, 63
Perkins, Michael, 78, 80, 81,'
89, 90, 102, 103, 105. 161-3,
166
Potts. F., 105
Procter, Joan, 84
Przibram, Doris, 122
Przibram, Gustav, 21, 22, 62
przibram, Hans, 21-4, 39, 42,
48, 49, 50, 60, 61, 62-3, 64,
68, 70-1, 72, 78, 86, 87, 98,
99-
100
, 102-3, 104, lOS.
108, 109, !I4, 115, 121, 123,
15
6
,
16
3, 164-5. 166-'7,
17
2 -3
Przibram, Karl, 21, !I4-I S
Przibram, Margeurite, 121
Przibram, Vera-see Countess
Teleki
Qu ASTEL, J. H., 71- 2 , 77-9,
80, 105. 166
REIFFENSTEIN, 123, 165, 166,
16
7
Romanes, George John, 57
Raux, 'Wilhelm, 22, 25, 26
ST,. HILAIRE, Geoffroy de, 130
Sahsbury, Frank B., 128, 130
Schnitzler, Arthur, 21
SchOnberg, Arnold, 20
Schulz, E., 172
Semon, Richard, 49
Shaw, George Bernard, 67, 81,
82
Smith, J. M., 128
Smythies, J. R., 132
Somerskjold, Grete-see Grete
Wiesenthal
Sonneborn, T. M., 130
Spallanzani, 152
Spetner, L. M., 128
Spiegelman, Sol.. 127
Steinach, Eugen, 22, 49
Stewart, B., 101, 123, 173
Swoboda, N., 141
TELEKI, Countess Vera, 49,
121-2
Temin, Howard, 127
Thorpe, W. H., 46, 76, 101,
105, 128, 164
Tower, W. L., 59
Tschermak vOn Seysenegg, E.,
53
UHLENHUTH, N., 101
VERMEER, J., ra7
Vevers, H. N., 105 .
Vries, H. de, 53
WADDINGTON, C. H., 30,
128, 130, 131-2
Wallace, 96
Walt, Anna, 94, 95
Walter, Bruno, 20
Watson, James D., 127
Watson, J. B., 92-3
Weismann, August, 29, 34, 37
Weiss, Paul, 22, 107--9, IIZ,
121, 123, 138
Werner, Franz, 99
Wettstein, R., 96-J
Wheldon, Muriel-----see the Han.
Mrs. Onslow
Wheldon, W. F. R., 54
Whyte, L. L., 130
Wiedersperg, Dr. Gustav von,
16,48
Wiesenthal, Bertha, 117
Wiesenthal, Elsa, 000
Wiesenthal, Grete, II7, II8,
II9, 138
Wiesenthal, Hilda, lI8
Wiesenthal, Martha, II7-18
About the Author
ARTHUR KOESTLER was born in 1905 in Budapest. Though he studied sci·
enee and psychology in Vienna, at the age of twenty he became a foreign
correspondent and worked for various European newspapers in the Mid-
dle East, Paris, Berlin, Russia and Spain. During the Spanish Civil War,
which he covered [rom the Republican side, he was captured and im-
prisoned for several months by the Nationalists, but was exchanged
after international protest. In 1939-40 he was interned in a French
detention camp_ After his release, due to British government interven·
tion, he joined the French Foreign Legion, subsequently escaped to
England and joined the British Anny.
Like many other intellectuals in the thirties, Koestler saw in the Soviet
experiment the only hope and alternative to fascism. He became a
member of the Communist Party in 1931, but left it in disillusionment
during the Moscow purges in 1938. His earlier books were mainly con·
cerned with these experiences, either in autobiographical form or in
essays or political novels. Among the latter, Darkness at Noon has been
translated into thirty languages.
After World War II. Mr. Koestler became a British citizen, and all
his books since 1940 have been written in English. He now lives in
London, but he frequently lectures at American universities, and was a
fellow at the Center for Advanced Study in the Behavioral Sciences at
Stanford in 1964-65.
In 1968 Mr, Koestler received the Sonning Prize at the University of
Copenhagen for his contributions to British culture. His works are now
being republished in a collected edition of twenty volumes.